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the wild yeasts were beneficial to growth<br />

under 12 sets of conditions tested. In yeasts,<br />

it has consequently now been established<br />

that prions are a naturally present<br />

supplementary source of inheritable material<br />

of adaptive value. The extent of prions in<br />

filamentous fungi as a whole has yet to be<br />

assessed, but they clearly have the potential<br />

to contribute to adaptability and fitness.<br />

volume 3 · no. 1<br />

Halfmann R, Jarosz DF, Jones SK, Change A,<br />

Lancaster AK, Lindquist S (2012) Prions<br />

are a common mehcnisms for phenotypic<br />

inheritance in wild yeasts. Nature 282:<br />

363–368.<br />

True HL, Lindquist SL (2000) A yeast prion<br />

provides a mechanism for genetic variation<br />

and phenotypic diversity. Nature 407:<br />

477–478.<br />

Different fungal and algal genotypes demonstrated<br />

within one lichen specimen<br />

Observations on the development of lichens<br />

in the field reveal that multiple propagules<br />

of a species developing on a surface often<br />

116<br />

P4T7e<br />

104<br />

P3T4c<br />

108<br />

P4T7<br />

102<br />

P3T4<br />

113<br />

P1T8<br />

103<br />

P5T3<br />

106<br />

P5T3c<br />

123<br />

P4T7d<br />

A single specimen of Parmotrema tinctorum showing the different fungal<br />

and algal genotypes determined with PCR of SSR markers. Codes prefixed<br />

by P are of the fungal partner, and those by T are of the algal partner; five<br />

fungal genotypes and one algal genotype were detected within this particular<br />

98<br />

specimen. P3T6 Adapted from Mansournia et al. (2012).<br />

Yeast colonies, light and tramsmission electron<br />

micrograph photos of Saccharomyces cerevisiae.<br />

coalesce to form a single structure. This is<br />

frequently observed where the propagules<br />

are asexual soredia or isidia, which may<br />

or may not have<br />

come from the same<br />

parent, and is welldocumented.<br />

However,<br />

whether all had to be<br />

of a single genotype<br />

for this to occur was<br />

uncertain. The first<br />

study to suggest that a<br />

single lichen specimen<br />

might not just have a<br />

single fungal partner<br />

experimentally was<br />

the study of Larson<br />

& Carey (1986) who<br />

found that single<br />

115<br />

P6<br />

Tabcde<br />

120<br />

P6<br />

Tabcde<br />

124<br />

P6<br />

Tabcde<br />

specimens of two<br />

Umbilicaria species<br />

showed variations<br />

in physiological<br />

parameters and<br />

isoenzyme profiles.<br />

With the advent<br />

of DNA PCR<br />

technology, and<br />

especially the use of<br />

microsatellite (SSR)<br />

markers, it has become<br />

possible to explore the<br />

<strong>issue</strong> of the degree of<br />

individuality of single<br />

lichen specimens<br />

with respect to both<br />

the fungal and the<br />

algal populations that<br />

comprise them.<br />

Parmotrema tinctorum is a rather<br />

common tropical lichen that reproduces<br />

mainly by asexual isidia. Mansornia et<br />

al. (2012) studied populations growing<br />

on Pinus thunbergii in Japan, and used<br />

microsatellite markers to characterize the<br />

partners at different levels: within single<br />

specimens, on single trees, and within 10 x<br />

10 cm quadrats. Of particular interest were<br />

the results from single specimens in which<br />

they studied numerous small pieces of t<strong>issue</strong>.<br />

They found that a single specimen could be<br />

formed from a single fungal partner with<br />

or without changes in the algal partner,<br />

or fusion of several independent partners.<br />

In total 12 fungal genotypes and 37 algal<br />

genotypes were recognized. An example in<br />

which there were five fungal genotypes and<br />

a single algal genotype is illustrated here.<br />

Further, specimens from individual trees<br />

or which were close together tended to<br />

have similar genotypes, suggesting limited<br />

dispersal in the site.<br />

This study provides evidence to support<br />

what has long been suspected, that one<br />

cannot presume that what looks like a single<br />

individual lichen specimen represents a<br />

single fungal genotype.<br />

Larson DW, Carey CK (1986) Phenotypic variation<br />

within “individual” lichen thalli. American<br />

Journal of Botany 73: 214–223.<br />

Mansournia MR, Wu B, Matsushita N, Hogetsu T<br />

(2012) Genotypic analysis of the foliose lichen<br />

Parmotrema tinctorum using microsatellite<br />

markers: association of mycobiont and<br />

photobiont, and their reproductive modes.<br />

Lichenologist 44: 419–440.<br />

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